Alien Terrestrial Arthropods of Europe, Part 2.

501 Pages·2010·23.6 MB·English

by  Alain Roques

Alien terrestrial
arthropods of Europe
Edited by
Alain ROQUES, Marc KENIS, David LEES,
Carlos LOPEZ-VAAMONDE, Wolfgang RABITSCH,
Jean-Yves RASPLUS and David B. ROY
Sofia–Moscow
2010
BioRisk 4(2) (Special Issue)
Alien terrestrial arthropods of Europe
Edited by Alain Roques, Marc Kenis, David Lees, Carlos Lopez-Vaamonde, Wolfgang Rabitsch,
Jean-Yves Rasplus And David B. Roy
Th is work was supported by a grant from the Sixth Framework Programme of the European
Commission under the project DAISIE (Delivering Alien Species Inventories in Europe), contract
SSPI-CT-2003-511202. We thank very much Jean-Marc Guehl (INRA department of «Ecologie
des Forêts, Prairies et Milieux Aquatiques») and Olivier Le Gall (INRA department of «Santé des
Plantes et Environnement ») for their fi nancial help which allowed to publish this book.
We are also very grateful to all colleagues who gently supplied us photos to illustrate the
alien species: Henri-Pierre Aberlenc, C. van Achterberg, Daniel Adam, G. Allegro, J.J. Argoud,
Margarita Auer, Juan Antonio Ávalos, Ab Baas, Antony Barber, Claude Bénassy, Christoph
Benisch, C. van den Berg, Mark Bond, Nicasio Brotons, Gert Brovad, Peter J. Bryant, David
Capaert, Jérôme Carletto, Rémi Coutin, David Crossley, Györgi Csóka, Massimiliano Di Gio-
vanni, Joyce Gross, L. Goudzwaard, Jan Havelka, Jean Haxaire, Franck Hérard, R. Hoare, R.
Kleukers, Zoltán Korsós, Gernot Kunz, Jørgen Lissner, Jean-Pierre Lyon, Mike Majerus†, Kiril
Makarov, Chris Malumphy, Erwin Mani, Paolo Mazzei, Tom Murray, Louis-Michel Nageilesen,
Laurence Ollivier, Jean-Pierre Onillon, Claude Pilon, Francesco Porcelli, Jean-Paul Raimbault,
Urs Rindlisbacher, Gaëlle Rouault, Gilles San Martin, R.H. Scheff rahn, Vaclav Skuhravý, John
I. Spicer, Massimo Vollaro, Jordan Wagenknecht, Beate Wermelinger, Alex Wild, Vassily Za-
khartchenko, and the Montpellier Station of the Laboratoire National de Protection des Végé-
taux, France. Olivier Denux did a great job in realizing all the distribution maps.
First published 2010
ISBN 978-954-642-555-3 (paperback)
Pensoft Publishers
Geo Milev Str. 13a, Sofi a 1111, Bulgaria
Fax: [phone]
[email protected]
www.pensoft.net
Printed in Bulgaria, July 2010
Contents
553 Chapter 10. Diptera
Marcela Skuhravá, Michel Martinez & Alain Roques
603 Chapter 11. Lepidoptera
Carlos Lopez-Vaamonde, David Agassiz, Sylvie Augustin, Jurate De Prins,
Willy De Prins, Stanislav Gomboc, Povilas Ivinskis, Ole Karsholt,
Athanasios Koutroumpas, Fotini Koutroumpa, Zdeněk Laštůvka, Eduardo Marabuto,
Elisenda Olivella, Lukasz Przybylowicz, Alain Roques, Nils Ryrholm,
Hana Šefrová, Peter Šima, Ian Sims, Sergey Sinev, Bjarne Skulev, Rumen Tomov,
Alberto Zilli & David Lees
669 Chapter 12. Hymenoptera
Jean-Yves Rasplus, Claire Villemant, Maria Rosa Santos Paiva, Gérard Delvare
& Alain Roques
767 Chapter 13.1. Thrips (Thysanoptera)
Philippe Reynaud
793 Chapter 13.2. Psocids (Psocoptera)
Nico Schneider
807 Chapter 13.3. Dictyoptera (Blattodea, Isoptera), Orthoptera,
Phasmatodea and Dermaptera
Jean-Yves Rasplus & Alain Roques
833 Chapter 13.4. Lice and Fleas (Phthiraptera and Siphonaptera)
Marc Kenis & Alain Roques
851 Chapter 13.5. Springtails and Silverfishes (Apterygota)
Jürg Zettel
855 Chapter 14. Factsheets for 80 representative alien species
Alain Roques & David Lees (Eds)
1023 Abbreviations and glossary of technical terms used in the book
Alain Roques & David Lees
1029 Index of the latin names of the arthropod species mentioned in the
book
Alain Roques & David Lees
BioRisk 4(2): 553–602 (2010) BA peerireovieweRd openiacscess jkournal
doi: 10.3897/biorisk.4.53 RESEARCH ARTICLE
www.pensoftonline.net/biorisk
Diptera
Chapter 10
Marcela Skuhravá1, Michel Martinez2, Alain Roques3
1 Bítovská 1227/9, 140 00 Praha 4, Czech Republic 2 INRA Centre de Biologie pour la Gestion des Popu-
lations (CBGP), Campus International de Baillarguet, 34988 Montferrier-sur-Lez, France 3 INRA UR633
Zoologie Forestière, 2163 Av. Pomme de pin, 45075 Orléans, France
Corresponding authors: Marcela Skuhravá ([email protected]), Michel Martinez ([email protected].
fr), Alain Roques ([email protected])
Academic editor: David Roy  |  Received 4 February 2010  |  Accepted 24 May 2010  |  Published 6 July 2010
Citation: Skuhravá M et al. (2010) Diptera. Chapter 10. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe.
BioRisk 4(2): 553–602. doi: 10.3897/biorisk.4.53
Abstract
Of the 19,400 native species and 125 families forming the European diptera fauna, 98 species (less than
0.5%) in 22 families are alien to Europe. Th ese aliens constitute 66 species (18 families) of the suborder
Brachycera and 32 species (4 families) of the suborder Nematocera. By family in this category, there are 23
Cecidomyiidae species, 18 Drosophilidae, nine Phoridae, eight Tachinidae and seven Culicidae. Another
32 fl y species belonging to fi ve families are considered to be alien in Europe. Th ese invasives native to
other European countries are composed of 14 species of Cecidomyiidae, seven Syrphidae, fi ve Culicidae
and three species each of Anthomyiidae and Tephritidae. Th e date of the fi rst record in Europe is known
for 84 alien species. Arrivals of alien species of Diptera have accelerated rapidly since the second half of
the 20th century. North America appears to be the dominant contributor of the alien fl ies. Th e majority of
alien Diptera were introduced into or within Europe unintentionally, with only three predators released
intentionally for biological control. Alien Diptera are predominantly phytophagous (35.6%), while a
lesser portion are zoophagous (28.6%) or detrivorous /mycetophagous (29.6%). Ecological impacts on
native fauna and fl ora have not been documented for any of the alien species established in Europe. How-
ever, 14 alien species have economic impacts on crops.
Keywords
alien, Europe, Diptera
Copyright Skuhravá M et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which
permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
554 Marcela Skuhravá, Michel Martinez & Alain Roques / BioRisk 4(2): 553–602 (2010)
10.1 Introduction
Diptera is one of the largest insect orders, with a worldwide distribution. Th e order
includes 172 to 179 families (depending on authors) with about 132,000 species de-
scribed which probably underestimates the actual fauna by at least a half. About 19,400
native species and 125 families have been recorded in Europe (Fauna Europaea). Th e
alien entomofauna is comparatively very limited with only 98 species observed to date,
i.e. less than 0.5% of the total dipteran fauna in Europe.
Commonly called true fl ies, mosquitoes, midges, deer- and horsefl ies and house-
fl ies feature among the most familiar Diptera. Flies are not only abundant in popu-
lar perception but also have particular veterinary and medical importance for vecto-
ring diseases and as pests of agriculture, forestry and husbandry. However, some spe-
cies are useful to man as parasitoids and predators of insect pests and as plant pollina-
tors. Generally, adults are minute to small, soft-bodied insects with a highly mobile
head, large compound eyes, antennae of variable size and structure, and sucking mou-
thparts. Th ey have only one pair of functional wings, the second pair being changed
into small head-like bodies called halteres. Legs are usually long, with fi ve-segmented
tarsi. Adults are usually very active and are found in all major habitats. Th ey are often
associated with fl owers and with decaying organic matter, but females of some groups
are blood-sucking. Larvae are eruciform and legless in most species. Th ey develop ma-
inly in moist or wet habitats such as soil, mud, decaying organic matter, and in plant
or animal tissues. Only a small proportion of larvae is truly aquatic. Th e majority are
liquid-feeders or microphagous.
10.2. Taxonomy of the Diptera species alien to Europe
Th e 98 species of Diptera alien to Europe belong to 22 diff erent families (Table 10.1),
which all have native representatives. A larger number of aliens belong to the sub-
order Brachycera (66 species and 18 families) than to the suborder Nematocera (32
species and 4 families). However, this apparently large diversity is confusing. More
than 40% of the alien species are either midges (Cecidomyiidae- 23 species) or fruit
fl ies and their relatives (Drosophilidae- 18 species). Th e other 20 families show less
than 10 species each (Figure 10.1). Th e arrival of these alien species has largely modi-
fi ed the composition of some families such as Braulidae and Drosophilidae where
at present aliens respectively account for 33.3% and 14.8% of the total fauna ob-
served in Europe. However, the native entomofauna includes 103 additional fami-
lies for which no alien species has yet been recorded in Europe, especially for some
ecologically and economically- important groups such as Chironomidae, Syrphidae,
Asilidae, Tipulidae and Anthomyiidae. Th e alien dipterans belong to the following
families:
Diptera. Chapter 10 555
Suborder Brachycera
Agromyzidae. All species in the family are phytophagous, including a number of seri-
ous pests of cultivated plants. Larvae live in plant tissues, usually forming characteristic
galleries as mines. Most larvae live in the parenchyma of leaves, or mine stems, few
attack fruits and seeds. Th e majority of the species are monophagous, some of them
are widely polyphagous, attacking diff erent plants of several families. To date, only fi ve
alien species have been observed in Europe relatively to 903 recorded native species
(Fauna Europaea). However, the alien fauna includes three species of Liriomyza (L.
chinensis, L. huidobrensis - see factsheet 14.23, and L. trifolii) which are highly damag-
ing to vegetable crops (Arzone 1979, Martinez 1982, Trouvé et al. 1991).
Braulidae. Larvae live as commensals within cells of honey-bee nests (Apis species).
Th ey feed on pollen, honey and organic debris. Adults are “food-parasites” of adult
bees, attaching themselves to the body of the queen or rarely to a worker. Th ey feed on
liquids from the mouth of the bees. Th ere is only one genus present in Europe, Braula,
which includes one alien species, B. schmitzi (Dobson 1999), and two native species.
Calliphoridae. Th is is a key family for human health. Adults are potential vectors
of bacteria, viruses, protozoaires and helminthes because they actively search for and
sit on feces, fresh and cooked meat, fi sh, dairy products, and wounds. Larvae are para-
sitoids or predators of living snails, or feed on blood of nestling birds. A few species
are obligate producers of myiasis in various animals. Only one alien species, Chrysomya
albiceps (Mercier 1927), has been observed in Europe compared to 112 native species.
Canacidae (=Tethinidae). Most species are strictly associated with salty habitats
(halobionts), e.g. coastal salt marshes, seashore wrack, sandy beaches, shores of inland
salt lakes, alkaline springs etc, and only a few species are also known from habitats
that are apparently without increased salinity (forests, meadows, deserts). Some species
have been reared from deposits of seaweed. Th ere is only one alien species, Pelomyia oc-
cidentalis (Irwin et al. 2001), compared to a total of 39 native species.
Ceratopogonidae. Biting adults of this family are potential vectors of major ani-
mal diseases. In particular, Culicoides species transmit bluetongue orbivirus between
ruminant hosts. A species of Afro-Asian origin, C. imicola Kieff er, has been conside-
red as the main agent of the recent outbreaks of bluetongue disease in Europe althou-
gh some native species could also be involved (e.g., C. pulicarius L. and C. newsteadi
Austen complexes (Purse et al. 2007)). However, it seems that the most likely mode of
incursion of C. imicola in Europe was via passive transport on the wind as aerial plank-
ton“ (Mellor et al. 2008, Purse et al. 2007). Th us, this species was not considered in
this chapter.
Dolichopodidae. Adults and larvae of most species are predaceous and feed on soft-
bodied invertebrates. Th ey occupy all terrestrial habitats from coastal beaches to high
elevations, but they generally prefer humid areas. Larvae are mostly found in moist so-
ils or in the litter layer while a few others depend on sap runs and tree rot holes for the-
ir development. Th ere is only one alien species, Micropygus vagans (Chandler 2004), in
comparison to 790 native species in Europe.
556 Marcela Skuhravá, Michel Martinez & Alain Roques / BioRisk 4(2): 553–602 (2010)
Figure 10.1. Relative importance of the families of Diptera in the alien and native entomofauna in Eu-
rope. Families are presented in a decreasing order based on the number of alien species. Species alien to
Europe include cryptogenic species. Only the most important families of native species (> 50 spp.) have
been considered. Th e number over each bar indicates the number of species observed per family.
Drosophilidae. Species in this family show very diverse biological habits. Th e lar-
vae of most species develop in fermenting substrates, but some mine living plants.
Some species are used as important laboratory animals. Drosophilids occur in all ter-
restrial habitats, from lowlands up to alpine meadows. Th ey may be found near the
habitats of their insect hosts or preys (mealybugs, bees, wood-boring beetles), around
toadstools (Polyporales) and in the fl ower heads of thistles. Aliens include 18 speci-
es in the genera Drosophila (8 species) (Bächli et al. 2002, Grassi et al. 2009), Chy-
momyza (4 species) (Band 1994, Carles-Tolra and Andersen 2002, Perju 1959, Trent
Band et al. 2005), Zaprionus (3 species) (Chassagnard and Kraaijeveld 1991, Monclus
1976, Tsacas et al. 1977), Scaptomyza (2 species) (Nicoli Aldini 2005, Nicoli Aldini
and Baviera 2002) and Dettopsomyia (1 species) (Prevosti 1976) compared to 104 na-
tive species.
Ephydridae. Adults are usually associated with moist substrates, especially shores,
marshes and wet meadows. Some develop in decomposing matter or excrement, other
are leaf miners or parasitoids. Aquatic and semiaquatic habitats are typical of the fami-
ly. A total of 335 native species occur in Europe with only three alien species - in the
genera Elephantinosoma, Placopsidella and Psilopa (Gatt and Ebejer 2003).
Diptera. Chapter 10 557
Fanniidae. Species inhabit forests, rarely open landscape and wetlands. Larvae are
generally saprophagous and mostly feed on decaying organic matter as human or ani-
mal faeces, decaying material in gardens, and rotting leaf litter. Some species have
been reared from fungi, others occur in bird nests, burrows of vertebrates, and nests
of social Hymenoptera. Th ere is only one alien species, Fannia pusio (Carles-Tolra and
Andersen 2002), compared to 82 native species.
Heleomyzidae. Larvae develop in sporocarps of fungi or live in association with my-
celia in forest soil, some are necrophagous or saprophagous. Th ere is only one alien spe-
cies, Prosopantrum fl avifrons (Ismay and Smith 1994) compared to 145 native species
Hippoboscidae. Adults are bloodsucking ectoparasites of birds and mammals. Fe-
males of all species are macrolarviparous, i.e. retaining the larva in the uterus to the
end of the third instar. Th ere is only one alien species, Crataerina melbae (Popov 1995),
compared to 29 native species.
Milichiidae. Larvae are saprophagous and develop in decaying vegetation, wood
detritus, in nests of birds, ants (myrmecophilous species) and of other social insects,
but also in excrements, carrion, dead insects and snails. Adults of some species are com-
mensals or kleptoparasites of predatory insects and spiders. Th ere are two alien species,
in the genus Desmometopa (Roháček (2006b)), compared to 41 native species.
Muscidae. Larvae develop in various kinds of decaying organic matter, often
showing facultative or even obligatory carnivorous behaviour. Larvae of some species
appear to be predaceous during their entire larval life. Adults feed on nectar or plant
sap, sometimes also on decaying liquids and some species are predaceous. Some spe-
cies are adapted to anthropogenically-altered ecosystems. Blood-sucking species are
of medical and veterinary importance, being vectors of some diseases. Th ere are two
alien species, the sorghum pest Athrerigona soccata (Vercambre et al. 2000), and a pre-
dator of house fl ies, Hydrotaea aenescens (Rozkošný 2006, Saccà 1964), compared to
585 native species.
Phoridae. Adults are found in all types of terrestrial habitats, particularly in forests
and meadows but also in steppe-like and xerothermic sites. Food preferences of lar-
vae appear to be remarkably diff erent. Most species are polysaprophagous with diff e-
rent degrees of specialisation. Parasitic species are often found in the nests of ants and
termites. Some fungus breeders feed on the fungi but others are obligate predators or
parasitoids of other fungus feeders such as larval Sciaridae. Th ere are nine alien species
in the genera Megaselia (three species) (Campobasso et al. 2004, Disney 2008, Disney
and Durska 1999), Chonocephalus (two species) (Disney 1980, Disney 2002), Dohr-
niphora (two species) (Disney 2002, Disney 2004), Hypocerides (one species) (Disney
2004), and Puliciphora (one species) (Disney 1983) in comparison to a total of 596
native species.
Sphaeroceridae. Larvae and adults are saprophagous. Larvae develop in diverse or-
ganic matter and feed as saprophages on microorganisms destroying rotting plants,
dung, carrion or fungi and also on the decomposed liquid substances. Adults occur in
all habitats that contain the breeding media of the larvae, preferably in damp places.
A few polyphagous species are synanthropic, living near human habitats. Many copro-